Revealing the significance of chlorophyll b in the moss Physcomitrium patens by knocking out two functional chlorophyllide a oxygenase.

The chlorophyllide a oxygenase (CAO) plays a crucial role in the biosynthesis of chlorophyll b (Chl b). In the moss Physcomitrium patens (P. patens), two distinct gene copies, PpCAO1 and PpCAO2, are present. In this study, we investigate the differential expression of these CAOs following light exposure after a period of darkness (24 h) and demonstrate that the accumulation of Chl b is only abolished when both genes are knocked out. In the ppcao1cao2 mutant, most of the antenna proteins associated with both photosystems (PS) I and II are absent. Despite of the existence of LHCSR proteins and zeaxanthin, the mutant exhibits minimal non-photochemical quenching (NPQ) capacity. Nevertheless, the ppcao1cao2 mutant retains a certain level of pseudo-cyclic electron transport to provide photoprotection for PSI. These findings shed light on the dual dependency of Chl b synthesis on two CAOs and highlight the distinct effects of Chl b deprival on PSI and PSII core complexes in P. patens, a model species for bryophytes.

Photoprotective Role of Neoxanthin in Plants and Algae.

Light is a paramount parameter driving photosynthesis. However, excessive irradiance leads to the formation of reactive oxygen species that cause cell damage and hamper the growth of photosynthetic organisms. Xanthophylls are key pigments involved in the photoprotective response of plants and algae to excessive light. Of particular relevance is the operation of xanthophyll cycles (XC) leading to the formation of de-epoxidized molecules with energy dissipating capacities. Neoxanthin, found in plants and algae in two different isomeric forms, is involved in the light stress response at different levels. This xanthophyll is not directly involved in XCs and the molecular mechanisms behind its photoprotective activity are yet to be fully resolved. This review comprehensively addresses the photoprotective role of 9'-cis-neoxanthin, the most abundant neoxanthin isomer, and one of the major xanthophyll components in plants' photosystems. The light-dependent accumulation of all-trans-neoxanthin in photosynthetic cells was identified exclusively in algae of the order Bryopsidales (Chlorophyta), that lack a functional XC. A putative photoprotective model involving all-trans-neoxanthin is discussed.

Mesophyll-specific phytochromes impact chlorophyll light-harvesting complexes (LHCs) and non-photochemical quenching.

Phytochromes regulate light-dependent plastid development and plant growth and development. Prior analyses demonstrated that phytochromes regulate expression of Sigma factor 2 (SIG2), which is involved in plastid transcription and coordinates expression of plastid- and nuclear-encoded genes involved in plastid development, as well as plant growth and development. Mutation of SIG2 impacts distinct aspects of photosynthesis, resulting in elevated levels of cyclic electron flow and nonphotochemical quenching (NPQ). As we initially identified SIG2 expression as misregulated in a line lacking phytochromes in mesophyll tissues (i.e., CAB3::pBVR lines), here we report on an investigation of whether photosynthetic parameters such as NPQ are also disrupted in CAB3::pBVR lines. We determined that a specific parameter of NPQ, i.e., energy-dependent quenching (qE) which is a rapidly induced photoprotective mechanism that dissipates stressful absorption of excess light energy during photosynthesis, is disrupted when mesophyll phytochromes are significantly depleted. The observed reduction in NPQ levels in strong CAB3::pBVR lines is associated with a reduction in the accumulation of Lhcb1 proteins and assembly or stability of light-harvesting complexes (LHCs), especially trimeric LHC. These results implicate mesophyll-localized phytochromes in a specific aspect of phytochrome-mediated NPQ, likely through regulation of chlorophyll synthesis and accumulation and the associated impacts on chlorophyll-protein complexes. This role is distinct from the impact of mesophyll phytochrome-dependent control of SIG2 and associated NPQ regulation.